Bird Conservation International (1995) 5:251-277 faunas of the humid montane forests of : biogeographic patterns and priorities for conservation

BLANCA E. HERNANDEZ-BANOS, A. TOWNSEND PETERSON, ADOLFO G. XAVARRO-SIGUENZA, and B. PATRICIA ESCALANTE-PLIEGO

Summary The distribution of 335 of in 33 islands of humid montane forest in Mesoamerica is summarized, and patterns of distribution, diversity and endemism are analysed. The montane forests of Costa Rica and western Panama far exceed other habitat islands considered for species-richness, richness of species endemic to Mesoamerica, and richness of species ecologically restricted to humid montane forests. Other , such as the Sierra Madre del Sur of and , the Los Tuxtlas of southern Veracruz and the mountains of and Guatemala, also hold rich and endemic avifaunas. Based on patterns of similarity of avifaunas, the region can be divided into seven regions holding distinctive avifaunas (Costa Rica and western Panama; northern and northern Chiapas; southern Chiapas; eastern north of the Isthmus of ; Sierra Madre del Sur; interior Oaxaca; and Transvolcanic Belt and ), which serve as useful guides for the setting of priorities for conservation action.

Se resumen las distribuciones de 335 especies de aves en 33 islas de bosque humedo de montana en Mesoamerica, y se analizan patrones de distribution, diversidad y endemismo. Los bosques montanos de Costa Rica y del oeste de Panama tienen la mas alta riqueza de especies, riqueza de especies endemicas a Mesoamerica, y riqueza de especies ecologicamente restringidas a bosque humedo de montana. Otras regiones, tales como la Sierra Madre del Sur de Guerrero y Oaxaca, la region de Los Tuxtlas y las montanas de Chiapas y Guatemala, tambien tienen avifaunas ricas en especies y en endemicas. Basado en patrones de similitud de avifaunas, se puede dividir Mesoamerica en siete regiones que tienen avifaunas distintas (Costa Rica y el oeste de Panama; el norte de Centroamerica y el norte de Chiapas; el sur de Chiapas; el este de Mexico; la Sierra Madre del Sur; el interior de Oaxaca; y el Eje Neovolcanico y la Sierra Madre Occidental), las cuales pueden servir como guias en el establecimiento de prioridades para la conservation.

Introduction Tropical forests are rapidly disappearing from the face of the earth, and with them are vanishing multitudes of species of plants and that depend on them for food and shelter. Reductions in forest cover in this century are frighteningly large, and have accelerated greatly in recent years owing to Blanca E. Herndndez-Banos et al. 252 expanding human populations, worsening economic situations, and improved technology for forest removal. For this reason, tropical forests have become the focus of intense interest from conservationists working to avoid large-scale loss of biological diversity (Sarukhan and Dirzo 1992). In spite of the interest in preserving tropical forest , the geographic distribution of that diversity is still largely unknown (Peterson et al. X993)- That is, for most taxonomic groups, very little is known about distributional limits of species, patterns of species-richness, or patterns of distribution of unique or endemic forms (Soule 1990, Escalante-Pliego et al. 1993, Peterson et al. 1993). Birds are an important group for such studies because their is well known, and because inventory methods are well developed, permitting detailed inferences from distributional data (e.g. ICBP 1992). Among the purposes of this paper, therefore, are to summarize the current knowledge of the distribution of birds in one habitat type - humid montane forests - in Mesoamerica, to analyse that information with regard to biogeographic patterns, and to begin to outline strategies for conservation action.

Humid montane forests in Mesoamerica For this paper, we define humid montane forest as middle-to-high elevation tropical forest that receives strong input of moisture from rain, fog, and clouds. This definition includes true cloud-forest, which is a dense mixture of temperate and tropical broad-leaved tree species with many epiphytes and tree ferns, as well as humid -oak forest. Tree species characteristic of northern Mesoamerican cloud-forests include members of the genera Quercus, Juglans, Podocarpus, and especially Liquidambar (east) and Chiranthodendron (west; Rzedowski 1986). Many genera and species of trees in eastern Mexican cloud-forests are also found in the eastern United States, probably indicating an ancient moist forest connection between the two regions (Martin and Harrell 1959). Humid pine-oak forests grade into true cloud-forests, and share with them structural features such as the abundance of tree ferns and epiphytes (Rzedowski 1986). Faunas of the two habitat types, when in contact, are almost identical (Navarro-Siglienza 1992, Torres-Chavez 1992), and therefore are considered together herein (Escalante-Pliego et al. 1993, Peterson et al. 1993). Humid montane forests are found in broken strings along both coastal slopes of Mesoamerica. This distribution makes intuitive sense: coastal mountains intercept moisture from clouds rolling in off the ocean. These forests are better developed in the east, where climatic conditions are more moist. The degree of development of cloud-forests in a particular region also depends on factors such as soil type and predominant weather patterns, but especially on the presence of high or steeply ascending mountains close to the coast. Hence, the geographic distribution of the habitat is everywhere fragmented and local (Figure 1). The dominant geographic features in Mexico are coastal ranges ("cordilleras") and interior basins. The western Cordillera consists of the Sierra Madre Occidental in the north-west, part of the Transvolcanic Belt from south to Michoacan, the Sierra Madre del Sur of Oaxaca and Guerrero, and the Sierra Madre de Chiapas. The eastern Cordillera includes the Sierra Madre Oriental in the north-east; part of the Transvolcanic Belt in Hidalgo, Puebla, and Veracruz; Bird faunas of Mesoamerican humid montane forests 253

\ c A,f ^7 .TAMPS S1NDU \ \ ^^^ HIDAL yy //^MALIN NAYJAt-V—\ /ACVERA '— v /-- r TAN C1—5v'~- . MICHO /\ Jtf/ GUACU

TEMAS y .NICAR YUCUN//T GU ERR / > ^T [ /CRGUA CUATR' KIAHU / P^f-^ /, CRTIL - ' //CRCEN // / //.CRTAL 1 ^ 1 KONDU ^ GUAPA 1 Figure I. Map of the distribution of patches of humid montane forest in Mesoamerica, showing those included in this study and several that are yet to be studied. the Nudo de Zempoaltepetl of Oaxaca; and the Sierra Norte de Chiapas. Separating these two mountain chains are arid interior basins including the Chihuahuan Desert, Valle de Tehuacan, Valle de Oaxaca, and the interior basins of Chiapas. Farther south, in southern Central America, the topography is often more simple, with one central cordillera and lowlands along both coasts. An east-west transect across Mexico at almost any latitude therefore crosses a remarkably consistent set of vegetation types. First, one encounters wet tropical habitats in the Atlantic coastal lowlands. On the coastal slope of the first range of mountains is a narrow band of cloud-forest, and above that a band of humid pine-oak forest: these two habitats are the focus of the present paper. At the highest elevations, alpine pine-fir forest is found. In the interior basins, dry pine-oak forest, desert, or deciduous tropical scrub prevails. Descending the western slope, the same sequence of habitats is found, from alpine pine-fir forest through humid pine-oak and cloud-forest down to lowland tropical forests. However, because climates of the western lowlands are often less humid than those of the eastern slope, the west-slope tropical forests rarely approach the lush, evergreen quality of the east slope, and indeed are in many cases wholly or partly deciduous. Much of the geography of habitats in the country can be understood on the basis of this simplified scheme. Distributions of vegetation types in Central America are similar, except that in many areas the dry interior basins are lacking, and humid montane forest is therefore more continuous (Long 1995). Blanca E. Herndndez-Banos et al. 254

Mesoamerican humid montane forest avifaunas The birds of humid montane forests in Mesoamerica represent a surprisingly diverse aggregation of species, and rank high among habitats in Mexico in total species-richness (Navarro-Sigiienza 1992, Escalante-Pliego et al. 1993, Peterson et al. 1993, Long 1995). Among the birds present are species commonly associated with lowland rainforests (e.g. Ornate Hawk-eagle Spizaetus ornatus, Ochre-bellied Flycatcher Mionectes oleagineus), as well as species typical of high-elevation, alpine forests (e.g. Sharp-shinned Hawk Accipiter striatus, Golden-crowned Kinglet Regulus satrapa). Also present, and perhaps most interesting, are many species and genera that are restricted to the humid montane forest zone, such as toucanets of the genus Aulacorhynchus, several rare and endangered guans (genera Penelopina, Oreophasis, and Chamaepetes), and jays of the genus . This variety suggests several ideas about why humid montane forests hold such diverse avifaunas (Escalante-Pliego et al. 1993). One possibility is that they represent a meeting ground between habitats, and their avifaunas largely consist of mixtures of both lowland and highland elements. A second view is that Mesoamerica as a whole represents an area of overlap between two faunas - those of North and - and that its richness results from the intermixture of the two geographically separated avifaunas. A final possibility is that substantial diversification has occurred in situ, and that the increased diversity of humid montane forests results from speciation and diversification of bird taxa in humid montane forests. It should be noted that these three explanations are not mutually exclusive, and that one, two, or all three may actually have played roles in the development of the humid montane forest avifauna. Hence, a second set of objectives for this study is to lay a basis for a more thorough understanding of the historical processes that generated the diverse humid montane forest avifauna. Detailed analyses of patterns of distribution, diversity, and endemism can reveal hints as to the history of the region and its biota. Later, combining these results with hypotheses of the history of particular taxa - phylogenetic hypotheses - detailed historical explanations can potentially be developed.

Methods We gathered data on the distribution of bird species in 33 isolated patches of humid montane forest in Mesoamerica (Figure 1), defined here as the region lying between the Rio Bravo (U.S.A.-Mexico border) and the Isthmus of Panama. Forest patches were chosen to represent as much as possible regions of continuous humid montane forest without major interruptions such as deep river valleys or desert basins. Some patches had to be eliminated from consideration owing to insufficient data; for example, the Sierra Mazateca of northern Oaxaca was not considered because nothing whatsoever was known of its avifauna (Binford 1989; results of surveys are currently in preparation). Obviously, because humid montane forest is by nature fragmented, our "patches" really represent systems of related patches of different sizes; badly Bird faunas ofMesoamerican humid montane forests 255 needed are surveys in tiny and remote humid montane forest patches. Because of the concentration of our field surveys in the northern part of the study area (Mexico), more detailed analysis was possible within Mexico than in the mountain systems in Central America; regardless, cloud-forest patches appear to be more continuous in Central America (ICBP 1992, Long 1995), making a coarser-grained view appropriate. Detailed analyses of the extent and condition of individual forest patches, as well as analyses of species-area relationships, must await detailed study of satellite imagery from the region. Three hundred and thirty-five bird species were included in the database for this study, based on the taxonomy of the American Ornithologists' Union (AOU 1983), with minor amendments to reflect more recent taxonomic opinion (Appendix 1). We used brackets [ ] to indicate the superspecies relationships of differentiated allopatric forms that would be considered semispecies (AOU 1983) or phylogenetic species (Cracraft 1983, McKitrick and Zink 1988). To restrict the analysis to taxa that actually breed in the study areas, we eliminated all species that occur only as migrants, vagrants, or winter residents (Remsen 1994). A few species, which have broad geographic distributions but occur in humid montane forests only marginally throughout much of their ranges, such as Scrub Jays coerulescens, are included in the data matrix only where they range broadly into that habitat. To focus the study on inhabitants of humid montane forests, we excluded species not living within the forest, for example, swallows that hunt for food above the canopy; families of birds that were thus eliminated include Hirundinidae and Alcedinidae. Based on descriptions of distributions (AOU 1983) and habitat use (see references), we described each species as occurring farther north (N = 1), farther south (S = 1), higher (A = 1), or lower (B = 1) in the altitudinal sequence of habitats. We define species occurring neither farther north nor farther south (N + S = o) as endemic to Mesoamerica, and species occurring neither higher nor lower in the altitudinal sequence of habitats (A + B = o) as ecologically restricted to humid montane forests. A wide variety of sources (e.g. Friedmann et al. 1950, Miller et al. 1957, Wetmore 1965-1984, AOU 1983, Stiles and Skutch 1989) was used to make the database reflect the known distributions of species as completely as possible. In addition, we consulted taxonomic treatments for many groups, faunal lists for numerous localities, and specimen data from several North American museum collections. For each definite record, a "1" indicating presence was entered in the appropriate cell of the data matrix; absences were entered as "o". Because, in many cases, existing inventories for particular montane forest habitat islands were incomplete, we also included a "probable" occurrence category: if a species occurred in habitat islands both north and south of an island in question, and if particuliar habitat requirements were met, then an "X" was entered in the appropriate cell of the data matrix. Later, we tested the validity of these distributional assumptions by alternatively setting all X = 1 (assumptions correct) or all X = 0 (assumptions incorrect), and reanalysing the data. A variety of approaches was employed to understand geographic patterns in the dataset. Species-richness and levels of endemism were assessed by simple counts of occurrences at each locality. Phenetic analyses of similarity among avifaunas of different localities were conducted by calculating Simpson's Index Blanca E. Herndndez-Baiios et al. 256 of Faunal Similarity, and then using the unweighted pair-group method of analysis (routines in SYSTAT, version 3.1) to produce dendrograms. Reanalysing the data under the two distributional assumptions mentioned above had no qualitative effect on patterns detected, and so only the results based on the most complete information (assumption X = 1) are presented herein.

Results

Patterns of species-richness and endemism Species-richness of birds in the humid montane forest islands in Mesoamerica snowed a broadly increasing trend from north to south (Table 1, Figure 2a). Forest islands in northern and central Mexico generally held 100 bird species or fewer, those in Chiapas and northern Central America 150-180 species, and those of Costa Rica and western Panama 180-201 species. Hence, the coarsest-scale trend was one of elevated species-richness in the southernmost montane forest islands. Exceptions to this trend included montane forest islands in the interior and those that are geographically isolated from the main mountain masses of Mesoamerica. For example, the forest islands in the interior of Mexico in the Transvolcanic Belt and central Oaxaca (Table 1, Figure 2a) held substantially fewer species than adjacent forest islands on the coastal slopes. Similarly, the isolated humid montane forests of the Los Tuxtlas region of south-eastern Veracruz held relatively few bird species (Andrle 1967). Surveys in small and isolated habitat islands in southern Mesoamerica would provide an important test of the generality of this result. In all, 145 species of birds that occur in humid montane forests are endemic to Mesoamerica (i.e. N + S = o). Patterns of species-richness among endemics of Mesoamerica were generally similar to patterns of species-richness in the region (Table 1, Figure 2b): northernmost forest islands contained few Mesoamerican endemics, whereas Central American forest islands, especially those of Costa Rica and western Panama, were extremely rich in such species. These patterns were apparent regardless of whether they were based on raw numbers of endemic species or on the percentage of endemism relative to the total species-richness at the site.

Patterns of distribution of habitat-restricted species A total of 147 species occur in Mesoamerica that are ecologically restricted to humid montane forests (i.e. A + B = o). These species are narrowly restricted to the humid forest band at middle elevations (1,000-1,800 m in Mexico). Geographic patterns of richness of these taxa were generally similar to those of species-richness: a broad trend of increasing numbers from north to south (Table 1, Figure 2c). Especially notable was the virtual absence of ecologically restricted species in the marginal northernmost forest islands in northern, central and western Mexico. Bird faunas of Mesoamerican humid montane forests 257

Figure 2. Distribution of species-richness of birds in the humid montane forests of Mesoamerica. Top: species-richness (white, 72-99; light grey, 100-136; dark grey, 145-178; cross-hatched, 195-200 species). Middle: richness of species endemic to Mesoamaerica (white, 3-11; light grey, 12-25; dark grey, 26-39; cross-hatched, 40-57 species). Bottom: richness of species restricted to humid montane forests (white, 7-15, light grey, 16-27; dark grey, 36-62; cross-hatched, 79-102 species). Blanca E. Herndndez-Banos et al. 258

Table 1. Summary of species-richness, richness of species ecologically restricted to humid montane forests, richness of species endemic to the humid montane forests of Mesoamerica, and richness of narrow endemics (at scales of 1-2 patches, 1-5 patches, and 1-10 patches), of 33 humid montane forest patches in Mesoamerica. Xo and Xi represent the two different assumptions made regarding completeness of inventories in each forest patch. Totals refer to numbers in each category in Mesoamerica as a whole. See Appendix 2 for patch names.

Species-richness Ecolog. restricted Endemic Narrow endemics

Patch Xo Xi Xo Xi Xo Xi 1-2 1-: ; 1-10 Eastern Mexico TAMPS 79 103 13 14 8 8 0 0 0 HIDAL 100 136 23 27 12 15 1 1 2 CVERA 139 175 36 41 20 23 1 2 4 TUXTL 82 104 18 24 11 14 2 2 3 CUATR 42 72 8 10 4 5 0 0 1 JUARE 125 154 34 38 20 22 0 1 3 FELIP 78 104 15 1.7 7 9 0 1 2 ZEMPO 71 154 21 36 12 20 0 1 3 SCRIS 81 165 36 51 27 36 0 1 14 MONTE 34 166 17 51 12 36 0 1 14

Western Mexico SINDU 36 95 4 10 3 4 1 1 1 NAYJA 71 122 12 17 9 9 1 1 2 MICHO 78 113 9 14 5 8 0 0 1 GUERR 111 133 25 27 15 15 1 2 3 YUCUN 68 103 15 16 6 6 1 2 2 MIAHU 99 124 23 24 12 12 1 2 3 CHIMA 97 120 32 36 21 22 0 0 4 SOCON 79 155 44 56 30 37 0 2 14 TACAN 65 156 36 57 27 38 0 2 15

Transvolcanic Belt,Mexico TANCI 58 93 8 11 4 5 0 0 0 TEMAS 57 91 7 11 4 6 0 0 1 TAXCO 54 80 7 7 4 4 0 0 0 MALIN 14 74 0 7 0 3 0 0 0

Central America GUAPA 145 178 52 58 36 38 0 3 16 GUACU 88 183 40 59 3° 39 0 2 16 GUAMI 95 181 38 58 26 38 0 1 15 HONDU 173 183 60 62 38 39 1 2 16 NICAR 91 147 26 43 20 23 1 2 6 CRGUA 130 145 53 58 26 28 0 18 20 CRTIL 159 170 74 79 38 40 0 28 30 CRCEN 193 195 96 97 54 54 0 41 43 CRTAL 190 195 97 99 56 56 3 43 45 WPANA 188 200 97 102 56 57 6 45 47

TOTALS 335 M7 101 M 61 78 Bird faunas of Mesoamerican humid montane forests 259

Of the ecologically restricted species, more than two-thirds (101 species) are also geographically restricted to Mesoamerica (i.e. N + S + A + B = o). This assemblage of species is found nowhere in the world but in Mesoamerican humid montane forests, and might be taken as a set of species that evolved in situ. These species are of special interest in conservation planning because of their extreme vulnerability to destruction of humid montane forests in a limited region.

Narrow endemics Species with extremely restricted distributions, "narrow endemics", are im- portant features in studies of biological diversity and endemism. Of the 335 species included in this study, 101 were endemic to Mesoamerican humid montane forests. Of these species, eight (7.9%) were restricted completely to a single forest island; 14 (13.9%) were found in one or two islands; and 61 (60.4%) were found in fewer than five islands of montane forest. Hence, extreme geographic restriction is common among the endemic birds of humid montane forests in Mesoamerica. The geographic distribution of these restricted-distribution species was focused in but a few zones (Figure 3). Species with geographic ranges of but one or two forest islands were concentrated in the Los Tuxtlas region of south- eastern Veracruz, the mountains of western Mexico, and especially in south- easternmost Costa Rica and western Panama. Each of these areas represents a mountain massif holding forests which were probably isolated from other humid montane forests since before the Pleistocene (Martin and Harrell 1959, Graham 1993), providing the isolation necessary for speciation to take place. Expanding the scale of endemism to include species with geographic ranges of five or fewer forest islands, the pattern was similar, although forest islands in north-eastern Mexico also appeared as hotspots, and the importance of the Costa Rican and Panamanian islands became much more emphasized. Expanding the scale still more, say, to ranges of up to 10 forest islands, emphas- ized Chiapas, Guatemala, Costa Rica and western Panama greatly, and de- emphasized the importance of forest islands farther north. Hence, the geo- graphic patterning of endemism is very much a function of the spatial scale at which endemism is defined.

Patterns of similarity among avifaunas Analyses of similarity among avifaunas of different forest islands (Figure 4) revealed clear geographic patterns. The avifaunas of the mountains of Costa Rica and western Panama were very different from those of forest islands farther to the north. The avifaunas of the forest islands of eastern Mexico north of the were all very similar to one another, and as a unit were most similar to those of Chiapas and northern Central America (within which the mountains of southern Chiapas segregated as a distinct entity). Three forest islands, the Chimalapas region of eastern Oaxaca, the Los Tuxtlas region of south-eastern Veracruz, and the mountains of Nicaragua, clustered loosely together, which, at least in the case of the latter island, makes no clear geo- Blanca E. Herndndez-Banos et al. 260

Figure 3. Distribution of richness of species of restricted geographic distribution (narrow endemics) in Mesoamerica, under different definitions of endemism. Top: 1-2 forest patches (white, o; light grey, 1; dark grey, 2; cross-hatched, 3-6 species). Middle: 1-5 forest patches (white, o; light grey, 1-3; dark grey, 18-28; cross-hatched, 41-45 species). Bottom: 1-10 forest patches (white, o; light grey, 1-6; dark grey, 14-20, cross-hatched, 30-47 species). Bird faunas of Mesoamerican humid montane forests 261

CRTAL CRTIL WPANA Southern Central Americ CRCEN CRGUA JUARE ZEMPO HI DAL Eastern Mexico (north ol TAMPS Isthmus) CVERA HONDU -, MONTE SCRIS GUAPA Chiapas and Northern GUAMI Central America GUACU SOCON TACAN CHIMA -i NICAR Poorly Surveyed Islands TUXTL TAXCO -• TEMAS MALIN TANCI Transvolcanic Belt MICHO NAYJA SINDU Sierra Madre Occidental GUERR -, MIAHU Sierra Madre del Sur YUCUN CUATR -i Interior Oaxaca FELIP Figure 4. Patterns of similarity among the avifaunas of patches of humid montane forest in Mesoamerica, based on UPGMA analyses of Simpson's Index of faunal similarity. graphic sense, although given their low level of similarity, they could perhaps be considered as three separate entities; perhaps they grouped together because their avifaunas are depauperate and/or poorly known. Finally, the forest islands of the Transvolcanic Belt and Sierra Madre Occidental of central and western Mexico (within which can be seen three geographic groupings: the Sierra Madre Occidental, the mountains along the coast from Michoacan to Nayarit, and the interior portion of the Transvolcanic Belt) and the mountains of southern Mexico west of the Isthmus of Tehuantepec (within which can be discerned a Sierra Madre del Sur group and an interior Oaxaca group) each formed units, and grouped together form a more inclusive unit separate from that of eastern Mexico and northern Central America. Hence, most forest islands included in Blanca E. Herndndez-Banos et al. 262

Figure 5. Biogeographic units based on patterns of similarity of avifaunas in the humid montane forests of Mesoamerica (numbers are equivalent to those on opposite page). this study fell into distinct geographic units, and biogeographic entities were thereby easily delineated (Figure 5).

Discussion Geographic patterns in distribution of species The avifauna of the humid montane forests of Mesoamerica is distributed unevenly. A general trend of increasing diversity from north to south across the study area was found, with the southernmost islands holding about twice as many species as the northernmost. Also, patches of humid montane forest isolated from the main Cordilleras had considerably fewer species than central patches. All of these patterns were perhaps expected given the ease of coloniza- tion of central as compared with peripheral forest patches, long-term stability and integrity of forest patches, patterns of complexity of plant communities, and the broad-scale trend of increasing diversity at lower latitudes. More interesting were the geographic patterns of distribution of species endemic to Mesoamerican humid montane forests. By far the greatest concentra- tion of endemic species in the region was in south-eastern Costa Rica and west- ern Panama. This result would seem surprising in view of the geographic prox- imity of the cloud-forests of eastern Panama and the of South America; however, as has been appreciated by other authors (e.g. Wetmore 1965-1984, ICBP 1992, Long 1995), many species have their geographic distributions Bird faunas of Mesoamerican humid montane forests 263 restricted to a tiny area very close to a barrier that separates the Mesoamerican and South American faunas. Such a concentration, with essentially no introgres- sion from forms to the east, suggests that the Isthmus of Panama has constituted an extremely strong barrier to dispersal, that colonization events in either direc- tion have been rare, and that connections of humid "montane" forest across the Isthmus were probably never established during the climatic fluctuations of the late Pleistocene glaciations.

Biogeographic boundaries The analyses presented herein, especially the cluster analyses of avifaunal simil- arity (Figure 4), allowed detailed and quantitative identification of biogeographic regions (Figure 5). The analyses presented above clearly identified seven geo- graphic clusters with characteristic avifaunas. In the list that follows, endemic species are listed, and selected species having distributions almost limited to the region are given in parentheses. (1) Costa Rica and western Panama: e.g. Pselliophorus spp., Pyrrhura hoffmani, Parula gutturalis and Troglodytes ochraceus, among others. (2) Northern Central America and northern Chiapas: Lampornis sybillae, Otus barbarus and melanocyaneus. (3) Southern Chiapas: (Tangara cabanisi) and (Oreophasis derbianus). (4) Eastern Mexico north of the Isthmus of Tehuantepec: Dendrortyx barbatus, (Cyanolyca nana) and (Rhynchopsitta terrisi). (5) Sierra Madre del Sur: Eupherusa poliocerca, E. cyanophrys, Cyanolyca mirabilis, (Nyctiphrynus mcleodii) and (Cypseloides stored). (6) Interior Oaxaca: (Cyanolyca nana). (7) Transvolcanic Belt and Sierra Madre Occidental: Cyanocorax dickeyi, Thalur- ania ridgwayi, (Streptoprocne semicollaris), (Rhynchopsitta pachyrhyncha), (Cardellina rubrifrons) and (Atlapetes virenticeps). Each of these regions is faunistically distinctive from other such regions. Of great interest to biogeographers is the historical pattern of interconnections among areas. This pattern could be represented as a tree showing the history of connection and isolation of different forest patches. The hierarchical pattern of similarity resulting from these analyses is as follows: (Costa Rica and western Panama, ((eastern Mexico, (northern Central America and Chiapas, southern Chiapas)), (Transvolcanic Belt and Sierra Madre Occidental, (Sierra Madre del Sur, interior Oaxaca)))) (Figure 4). Although this pattern was detected by phenetic analyses, to the degree that avifaunal similarity reflects historical rela- tionship, we can take it as a preliminary hypothesis of historical relationships among cloud-forest areas for later testing based on phylogenetic studies.

Endemism and speciation By inspecting patterns of distribution of endemic species, regions that may be foci of differentiation and speciation may be identified. For example, the Costa Rica and western Panama area has many endemic species that appear to have Blanca E. Hernandez-Banos et al. 264 differentiated and speciated in situ. Other areas that represent potential foci of differentiation and speciation include the Los Tuxtlas region of south-eastern Veracruz and the Sierra Madre del Sur of Guerrero and Oaxaca. Detailed phylo- genetic studies of individual taxa are needed to clarify the historical processes that underlie these distributional patterns. Although the high endemism in the Costa Rica/Panama area has been well appreciated (Wetmore 1965-1984, Stiles and Skutch 1989, ICBP 1992), elevated endemism in the two Mexican regions has gone relatively underappreciated (but see ICBP 1992, Long 1995). In 1988-1989, when three of us wrote a summary of patterns of avian diversity in Mexico (Escalante-Pliego et al. 1993), we knew of but one species endemic to the Los Tuxtlas region, the Campy- lopterus excellens. Since that time, however, we have come to appreciate the distinctiveness of three other taxa: the quail-dove Geotrygon carrikeri, which is recognizable as a species separate from the southern G. lawrencii based on char- acters of plumage and morphology (Peterson 1993); the bush-tanager Chloro- spingus [ophthalmicus] wetmorei, which is distinctive in genetic characters (Peterson et al. 1992); and the brush-finch Atlapetes [brunneinucha] apertus, which is readily distinguishable on the basis of plumage characters. Recognition of these forms as separate species of course depends on the species concept used; nevertheless, they are diagnosable forms and represent units of biological diversity distinct from other populations currently placed in the same species. Hence, the Los Tuxtlas region actually holds a number of endemic taxa, although the first impression was to the contrary; assessment of its importance as a centre of endemism if all near-species-level differentiates were recognized, however, must await future analyses (Navarro-Sigiienza and Peterson in prep.). The Sierra Madre del Sur of Guerrero and Oaxaca (in some taxa extending north-west to Michoacan, Jalisco and Nayarit) is still more striking in its richness of endemic species, the distinctiveness of many of which has also been underap- preciated. Taxa occurring in humid montane forests in the region include the newly described swift Cypseloides storeri (Navarro-Sigiienza et al. 1992), the hermit hummingbird Phaethornis [superciliosus] tnexicanus (R. C. Banks pers. comm.), the extremely rare hummingbird Lophornis brachylopha (Banks 1990), the Thalurania ridgwayi (Jalisco and Nayarit only: Escalante-Pliego and Peterson 1992), Eupherusa poliocerca*, E. cyanophrys*', and Lampornis [ameth- ystinus] margaritae (Torres-Chavez et al. in prep.), the toucanet Aulacorhynchus [prasinus] wagleri (Benitez-Diaz et al. in prep.), the woodpecker Piculus auricu- laris* (Baptista 1978), the antpitta Grallaria [guatimalensis] ochraceiventris (Peterson et al. in prep.), the jays Cyanolyca mirabilis* and Aphelocoma [unicolor] guerrerensis (Peterson 1992), and the bush-tanager Chlorospingus [ophthalmicus] albifrons (asterisks indicate recognition by AOU 1983). Hence, of the 13 species or near- species-level differentiates in humid montane forests in the region, only four were fully recognized by the AOU (1983). The majority are presently under study by ourselves and our students, based on a variety of morphological char- acters, investigations that are revealing more striking levels of endemism than previously appreciated (Peterson and Navarro-Sigiienza in prep.). The import- ance of the mountains of western Mexico as a centre of avian endemism has also been pointed out by ICBP (1992) and Long (1995). Viewed more generally, in the context of the question posed in the Introduc- tion, the surprising biological diversity of the humid montane forests of Bird faunas of Mesoamerican humid montane forests 265

Mesoamerica has a complicated history. Some taxa have clearly entered the region from north or south (see examples in Introduction), but fully 101 of the 335 species (30.1%) seem to have speciated in situ in Mesoamerican humid montane forests. Hence, all three general explanations may have played import- ant roles in the diversification of the birds of the humid montane forests of Mesoamerica (Escalante-Pliego et al. 1993).

Priorities for conservation action An important reason for conducting studies such as this is to aid the design of optimal strategies for conserving biological diversity. In the present case, the avifaunas of humid montane forests are in trouble because the habitat is increas- ingly endangered. Humid montane forests are cleared for lumber and paper pulp interests, planting coffee, corn, and other agricultural products, or for grazing cattle or goats. In some cases, the destruction is absolute (e.g. clear- cutting of forests); in other cases, it is gradual, such as through the grazing of cattle within forests, which causes a slow degradation of forest quality. The end result, however, is much the same: a place uninhabitable by animals and plants requiring native forest cover. Given that money and opportunities for conservation action are limited, care- ful design of a strategy for conserving maximally the biological diversity of these forests is critical. A common approach is to focus on species of particular inter- est: for the present study, candidates might include the Horned Guan Oreophasis derbianus and Resplendent Quetzal Pharomachrus mocinno, both of which are large, attractive, endangered, and endemic to Mesoamerican humid montane forest. Nevertheless, as we have demonstrated in these analyses, the set of species potentially threatened in Mesoamerican humid montane forests is much more inclusive, and we therefore believe that efforts to preserve maximally the biological diversity of the region must instead be focused at the level of communities. An exciting result that has emerged in the past decade is the remarkable coincidence in patterns of diversity and endemism among different taxonomic groups in Mesoamerica (see, e.g., Peterson et al. 1993, Ramamoorthy et al. 1993, Vazquez-Garcia 1995). Patterns being concordant among many taxa, optimal conservation strategies should also generally coincide, because endemic forms are concentrated in the same areas. Given that birds are such a well-known group in terms of distribution and diversity, analyses of patterns of avian divers- ity are critical to the establishment of conservation priorities. The biogeographic regions delimited above are a good first step. Viable reserves should be estab- lished in each of the seven regions: (1) Costa Rica and western Panama, (2) northern Central America and northern Chiapas, (3) southern Chiapas, (4) east- ern Mexico north of the Isthmus of Tehuantepec, (5) Sierra Madre del Sur, (6) interior Oaxaca, and (7) Transvolcanic Belt and Sierra Madre Occidental. Preservation of examples of each of these regions would protect the great major- ity of endemic montane forest birds in Mesoamerica. Unfortunately, not all seven areas are currently under protection. Those areas that are currently protected more or less adequately (i.e. excluding areas under decree as protected areas but not in actuality receiving any protection) are (1) Blanca E. Herndndez-Banos et al. 266

Costa Rica and western Panama (e.g. Parque Nacional La Amistad), (2) northern Central America and northern Chiapas (e.g. Parque Nacional Lagunas de Montebello), (3) southern Chiapas (Reserva de la Biosfera El Triunfo), and (7) Transvolcanic Belt (e.g. Parque Nacional Pico de Tancitaro, Parque Nacional Lagunas de Zempoala) and Sierra Madre Occidental (e.g. Reserva de la Biosfera "Sierra de Manantlan"). The remaining areas - (4), (5) and (6) - remain com- pletely unprotected. We list (4) as unprotected in spite of the existence of the Reserva de la Biosfera "El Cielo" in Tamaulipas, because that mountain range is too far north to hold many of the species characteristic of that region. Focusing on the 101 bird species endemic to Mesoamerican humid montane forests, the existing reserve system misses completely seven of these species: Cyanocorax dickeyi of the Sierra Madre Occidental of Sinaloa and Durango; the Bearded Wood-partridge Dendrortyx barbatus of Hidalgo, San Luis Potosi and northern Puebla and Veracruz; the Dwarf Cyanolyca nana presently restricted to Cerro San Felipe of central Oaxaca; the Blue-capped Hummingbird Eupherusa cyanophrys of the Sierra de Miahuatlan of southern Oaxaca; the White-tailed Hummingbird E. poliocerca; the White-throated Jay Cyanolyca mirabilis of the Sierra Madre del Sur of Guerrero and Oaxaca; and the Green-breasted Moun- tain-gem Lampornis sybillae of Nicaragua and Honduras. Five of these species - Dendrortyx barbatus, Cyanolyca nana, C. mirabilis, Eupherusa cyanophrys, and E. poliocerca - have already been identified as threatened with global (Collar et al. 1992). Hence, clear priorities in the situation of new reserves include these areas, especially the Sierra Madre del Sur of Guerrero and Oaxaca, where three of these species remain unprotected (not to mention 10 additional forms presently under study, as mentioned above). An ideal, if perhaps impractical, plan for the conservation of humid montane forest avifaunas would be the establishment of protected areas in each forest island, given that avifaunas of different islands within regions, although similar, are not identical. For areas in which political instability is a concern, binational protected areas may potentially buffer unique habitats against the whims of human matters (consider, for example, the destruction of forest areas in north- ern Chiapas in 1994 and 1995). More than anything, conservation of montane forest areas in Mesoamerica must adopt a two-pronged approach: (1) emergency protection of even the smallest patches of forest in critically endangered areas (e.g. forests in north-eastern Mexico, which are threatened with total eradication), and (2) identification and protection of the largest and most com- plete areas, which may permit the long-term preservation of entire ecosystems without substantial losses due to area effects (e.g. Sierra de Juarez, Oaxaca, where 300,000+ ha of montane forest remain almost untouched). Our hope is that information and analyses presented herein will serve to focus such efforts more precisely, and allow conservation planners to make optimal use of their resources. We can make available electronic copies of the data matrix so as to permit further analysis and interpretation of the information presented herein.

Acknowledgements We thank a great many people for their help with this project over the past 14 years. First and foremost, thanks to our many students who in the course of Bird faunas of Mesoamerican humid montane forests 267 their studies have dedicated hundreds of hours to the work in field, library, museum, and laboratory needed to assemble this database. We thank our dear friends, Claudia Abad de Navarro, Carlos Cordero, Livia Leon-Paniagua, and Rosa M. Salazar for their friendship, help, and encouragement during this pro- ject. Several colleagues have offered helpful comments on the manuscript, including Mark Robbins, Oliver Komar and an anonymous reviewer. For access to specimens, we thank the curators of the Field Museum of Natural History, American Museum of Natural History, and University of Michigan Museum of Zoology. Funding was provided by the National Geographic Society, National Science Foundation, Direccion General de Asuntos del Personal Academico, CONACyT, PADEP, and the Field Museum of Natural History.

Appendix 1. Summary of known distributions of birds in patches of humid montane forest in Mesoamerica. For each patch, occurrence of species is indicated as o, absent; x, not definitely known to occur, but likely on biogeographic grounds; and 1, definitely occurs. "HAB" = habitat use: A, occurs in habitats above humid montane forests; and B, occurs in habitats below humid montane forests. "GEO" = extent of geo- graphic range: N, occurs farther north than the northern limit of humid montane tropical forests as defined for the purposes of this paper; and S, occurs south or east of the Isthmus of Panama. "TV Belt" = the Transvolcanic Belt of central Mexico. Forest islands are numbered as in Appendix 2. Mexico HAB GEO East slope West slope TV Belt Central America A B N s

Abeillia abeillei ) 0 0 0 0 0 1 0 0 1 0 X 1 X 0 0 0 0 0 0 1 1 X 0 0 D 0 1 X X 1 10 0 0 0 0 Acanthidops bairdii ) 0 0 0 0 0 0 0 0 0 0 0 0 0 000000000 0 0 J 0 0 0 0 0 0 0 1 1 1 1 Accipiter bicolor 3 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 O 0 ) 0 1 X X X X 1 1 1 I 1

Accipiter gentilis 1 0 1 0 O 1 0 o 0 0 0 0 0 0 10 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 O 0 0 O 0 0

Accipiter striatus 0 1 1 X 10 0 0 0 10 0 X 0 10 1 0 0 1 1 1 10 0 0 1 1 1 1 1 0 0 0 0 0

Aegolius acadicus I 0 1 0 0 0 1 () 0 0 1 X0 0 0X00 0 0 1 0 0 10 0 1 0 0 0 0 0 0 0 0 0 0 Aegolius ridgivayi 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 X X 1 0 0 0 11 1 Aeronautes saxatalis 0 1 0 X 1X 0 0 0 0 0 0 0 X 0 1 10 0 0 0 I 1 0 0 1 X X 1 0 0 0 0 0 0

Amaurospiza concolor ") () 0 1 0 0 o 0 0 0 0 X 0 0 0 10 1 0 1 0 1 0 0 0 0 0 0 0 0 1 11111 1

Amazilia ben/llina 1 0 0 0 0 1 X 1 0 1 1 1 X X 1 1 1 X 1 0 X X 1 1 X 1 X 1 1 0 0 0 0 0 0

Amazilia Candida ) 1 o 0 0 1 1 1 0 1 0 X 1 1 0 0 0 0 0 0 X 0 0 0 0 J 0 1 X X 1 0 0 0 0 0 0

Amazilia cyanocephala 1 o 0 1 1 1 X 0 1 1 1 1 X 0 0 0 0 0 0 1 X X 0 0 J 0 1 1 1 1 10 0 0 O 0

Amazilia tzacatl ) 1 o 1 1 XX 1 X 1 X X X X 0 0 0 0 0 0 X X X 0 0 D 0 1 X X 1 11111 1

Amazona finschi ) 1 0 o 0 0 o 0 0 0 0 0 0 0 X X X X 1 1 0 0 0 0 0 D 0 0 0 0 0 0 0 0 0 0 0 Anabacerthia variegaticeps J 1 0 1 O 0 0 1 0 10 1 X 1 0 O 0 1 X 1 1 1 1 0 0 1 0 1 1 1 1 X 1 1 I 1 1

Aphelocoma coerulescens 0 1 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 O 0 0 Aphelocoma ultramarma 0 1 0 1 I 0 0 0 0 0 0 0 0 0 10 0 0 0 0 0 0 1 1 X 000000000 0 Aphelocoma unicolor _> 0 0 0 O 1 1 a 0 1 O 1 1 1 O 0 0 1 O 0 0 1 1 0 0 •) 0 1 1 1 1 0 0 O O 0 0 Aratinga liolociilora 0 1 0 1 1 1 X X XXX X X X X X X X X 0 0 0 0 0 ) 0 1 X X 1 1 0 O 0 0 0

Aratinga strenua 1 1 0 o 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 X X 0 O ) 0 1 0 0 1 X 0 0 0 0 0 Asio stygius Aspathn gularis 1 0 0 0 0 0 0 0 0 0 0 0 1 X 0 0 0 0 0 0 1 1 1 0 0 ") 0 1 1 1 I 0 0 0 0 0 0

Atlapetcs albinucha ) O 0 0 0 X 1 0 0 1 0 1 1 1 0 0000000 0 0 0 "1 0 0 0 0 0 0 O O 0 0 o

Atlapetes brunneinucha ) 0 0 1 1 1 1 1 1 1 1 1 1 1 0 0 0 1 1 1 11 1 0 0 D 0 1 1 1 1 X 1 1 1 1 1 Atlapetes gutturalis J 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 O 0 ) O 1 1 1 1 X 1 1 1 1 1 Atlapetes pileatus 0 I 0 1 1 1 0 X X "1 1 0 0 1111 X 1 0 0 0 1 1 X 0 0 0 0 0 0 0 0 0 0 Atlapetes virenticeps 0 I 0 O 0 0 0 0 0 O O 0 0 XI 1 0 O 0 0 0 0 1 1 X o 0 0 0 0 0 0 0 0 0 Atthis elhoti ) 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 X 0 0 0 0 1 1 X 1 0 0 0 0 0 0 Atthis heloisa o 0 0 1 X1 0 1 1 1 10 0 X 1 1 1 1 1 O0 0 X 1 X 0 0 0 0 0 0 0 0 0 0

Attila spadiccus 0 1 0 0 1 1 0 X 0 X X X Blanca Blanca E. Herndndez-Banos et al. 268

Appendix 1 (cont.) Mexico HAB GEO East slope West slope TV Belt Central America AB N S

Aulacorhynchus prasinus O111O1OX1X OOlllllll OOOO 1111111111 Automolus ochralaemus 001 1 x 1 x x x x 000000000 0000 1XX1111111 Automolus rubiginosus o x 1 0 0 1 0 i x x 0 00111011 OOOO 1111X0001 1

Basileuterus belli 1 11 1 0 1 1 1 1 X 1 11 1 1 1 1 1 X 1 1 0X 1 1 1 1 0 0 0 0 0 0 Bnsileuterus culicivorus 1 11 1 0 1 0 1 X X 0 1X 1 1 1 1 X X 0 0 0 0 1 X X I X 1 1 1 1 1 Basileuterus melanogenys 0000000000 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 Basileuterus tristriatus 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1

Bolborhynchus lincola 0 0 1 0 0 1 0 X 1 X 0 a » X 0 0 1 X 0 0 0 0 1 X X 1 X 1 1 1 1 1 Buteo jamaicensis

Campephilus guatimalensis 1 11 1 0 1 0 X 1 X X X 1 1 X X 1 X X 1 0 0 0 1 X X 1 1 1 1 1 1 1 Campybpterus curvipennis 1 11 0 0 1 0 1 X X 0 0 0 0 0 0 0 0 0 0 0 0 0 X X X 1 0 0 0 0 0 0 Campybpterus excellens 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Campylopterus hemilcucurus 0 0 1 1 0 1 0 1 X X 0 0 0 1 0 0 1 1 1 0 0 0 0 1 1 I 1 I 1 1 1 1 1 Campybpterus rufus 0 0 0 0 0 0 0 0 X X 0 0 0 0 0 0 1 1 X 0 0 0 0 1 X X 1 0 0 0 0 0 0 Campylorhamphus pusillus 0 0 0 0 0 0 0 0 0 0 0 0 000 0 O 0 G 0 0 0 0 0 0 0 0 0 0 1 1 1 1 Campylorhynchus rnegalopterus 0 0 1 0 0 1 1 1 0 0 0 1 1 0 0 0 0 0 0 1 1 10 0 0 0 0 0 0 0 0 0 0 Cajnpylorhynchus zonatus 0 1 1 1 0 1 0 X 1 1 0 0 a 0 0 0 1 X 1 0 0 00 I 1 I 1 1 1 1 1 1 1 Caprimulgus ridgzvayi 0 0 00 0 0 0 0 0 0 X 1 X X X 1 X X X X 0 0 0 X X X 1 X 0 0 0 0 0 Caprimulgus saturatus 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 Caprimulgus vodferus X X 1 X 1 1 1 X X 1 X 1 1 1 1 1 1 X X 1 1 1X 1 1 1 1 0 0 00 0 0 Cardellina rubrifrons 0 0 0 0 0 0 0 0 0 0 1 00 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Carduelis atriceps 0 0 00 0 0 0 0 1 X 0 0 0 0 0 0 0X X 0 0 0 0 1 X 0 0 0 0 0 0 0 Carduelis notata 1 11 0 1 0 1 1 X X 1 11 1 1 1 1 X 1 1 1 1X 1 1 1 1 I 0 0 0 0 0

Carduelis pinus 0 X 1 0 0 0 0 0 1 X X X 0 0 0 0 0 0 0 X X 0 1 0 X X 0 0 0 0 0 0 0

Carduelis xanthogastra 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (1 1 1 1

Catharus aurantiirostris X 1 1 0 0 1 1 1 X X 1 11 1 1 1 I X 1 X 1 XX 1 X X 1 1 X 1 1 0 1

Catharus dryas 0 0 00000000 0 0 0 0 0 0 1 1 1 0 0 0 0 1 1 1 1 0 0 0 0 0 0 Catharus frantzii 0 0 1 0 0 1 1 1 1 X 0 1 0 I 0 I 1 1 I 1 0 0 0 1 1 1 1 X 1 1 1 1 1 Catharus fuscater 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1

Catharus gracilirostris 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 1 1 1 Catharus mexicanus 1 11 1 0 1 0 0 1 X 0 0 0 0 0 0 1 X X 0 0 0 0 X 1 1 1 X I 1 I 1 1

Catharus occidentalis 0 0 1 0 0 I 1 1 0 » 0 1 1 1 1 1 0 0 0 1 1 1X 0 0 0 0 0 <> 0 0 0 0 Catherpes mexicanus

Certhia familiaris X X X <> 1 1 1 X 1 X 1 X1 1 0 1 <> X 1 1 X X 1 1 1 1 1 1 0 0 0 0 0 Chaetura vauxi

Chamaepetes unkolor 0 0 0 0 0 0000 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 I Chlorophonia callophrys 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 1 1 1 1 1 Chlorophonia occipitalis 0 0 1 1 0 1 0 X 1 X 0 0 0 0 0 0 1 1 1 0 0 0 0 1 X X 1 1 0 0 0 0 0

Chlorospingus canigularis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (1 0 0 o 0 0 0 0 0 0 0 0 1 0 1 Chlorospingus ophthalmic us 0 1 1 1 0 1 0 1 1 1 0 0 0 1 I 1 1 I 1 0 0 0 0 1 1 1 1 X 1 1 1 1 1

Chlorospingus pileatus 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 » 0 0 1 1 1 Ciccaba virgata

Cinclus mexicanus 0 0 1 0 0 0 1 X 0 0 X X X 1 0 1 0 0 0 1 1 00 1 1 X 1 I 1 1 1 1 1

Claravis mondetoura 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 I X 1 1 X 1 1 1 1 1

Coccothraustes abeillei 1 11 X 0 1 0 X X 1 X 1 X I 0 0 0 I ] X X 1 0 1 1 1 1 0 0 0 0 0 0

Coccothraustes vespertinus 0 X 1 0 0 0 1 0 0 0 X X X 0 0 0 0 0 0 X X 0 0 0 0 0 0 0 0 0 0 0 0

Colaptes auratus X 1 I X 1 1 1 1 1 X X X I 1 1 1 X X X 1 X 1 X ] I 1 I 0 0 0 0 0 0

Colibri delphinae 0 0 0 0 0 0 o 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 X 1 X 1 1 1 1 1 Colibri thalassinus 0 1 X 0 1 1 1 1 1 X 0 X 1 1 0 1 1 1 X 1 1 10 1 1 X 1 X X I 1 1 1

Columba fasciata X X X 0 1 1 1 X 1 X X 1 1 1 1 1 1 1 X X X X X 1 1 1 1 1 1 1 1 1 1 Columba subvinacea 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 (1 0 0 0 0 0 0 0 0 0 0 1 I 1 1 Contopus lugubris 000 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 Contopus ochraceus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 Contopus pertinax 1 11 0 1 1 1 X X X 1 11 1 I 1 1 X X 1 1 XX ] 1 1 1 1 0 0 0 0 Cotinga amabilis 0 0 0 1 1 0 X X X 0 0 0 0 00 0 0 0 0 0 0 0 X X \ 1 X 1 1 1 1 0 Cotinga ridgwayi 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 Cranioleuca erythrops 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 Crax rubra X X X 1 X 1X X X X X (1 0 X X X 1 1 I I 1 1 Bird faunas of Mesoamerican humid montane forests 269

Appendix 1 (cont.) Mexico HAB GEO East slope West slope TV Belt Central America A B N S

Crypturellus cinnamomeus 1 O O X X X X 1XX1XX1 IX 0000 1 X X 1110 0 0 1 Cyanerpes cyaneus 000000000 OOOO 1X11 1 1 1 1 1 X stelleri 0 > 10 0 0 0 0 Cyanocompsa parellina 1 1 17 0 0O 1 X X XXX X 1 1 1 X 1 0 0 0 0 XXXI 100000 Cyanocorax dickeyi 0 0 00 0 00 0 0 0 10 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 Cyanocorax melanocyaneus 0 0 00 o o 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 1X11 10 0 0 0 0 Cyanocarax yncas 111 1 O X 0X11 O 1 11 X 1 1 1 X 0 0 0 0 1 X X 10 0 0 0 0 0 Cyanolyca argentigula o o 00 0 00000 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 111 Cyanolyca cucullata Oil 0 0 10 111 0 0 00 0 0 111 0 0 0 0 1111 10 1111 Cyanolyca mirabilis o o o0 0 00000 0 0 01110 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 Cyanolyca nana 0 0 10 0 1110 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 Cyanohjca pumilo 0 0 00 0 00 0 11 0 0 00 0 0 0 1 1 0 0 0 0 1111 0 0 0 0 0 0 Cydarhis gujauensis 1 1 11 O X 0 X 1 X 0 0 0O O 0 1 X X 0 0 0 0 1 X X 1 1X1111 Cypseloides chcrriei 0 0 00 0 00 0 0 0 000 0 0 0 0 0 0 0000 0000 o 0 o 1 1 x Cypseloides cryptus o o o0 0 00 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 O O X 0 1 1 1 1 X Cypseloides niger 0 0 10 0 110 0 0 XXX 0 0 1 0 X X X 0 0 X XXXI XXXI IX Cypseloides rutilus 0 0 1X 0 10 1 1 X X 1 00 1 i 1 x x 0X00 XXXI X X X 1 1 1 Cypseloides storeri 0 0 0000 0 0 0 0 0 0 010 0 0 0 0 X O O 0 0000 000000

Dadylortyx thoracicus 1 X 1X X X X X 1 X O 1 X1 0 0 1 1 X 0 0 0 0 1 1 X10 0 0 0 0 0 Dendrocincla anabatina O 1 X1 0 1 O X X X 0 0 00 0 0 0 0 0 0 0 0 0 XXI 110 X X 1 1 Dcndrocinda homochroa 000 0 0 1 O 1 X X 0 0 00 0 O 1 X X 0 0 0 0 1 X X 1X1 X X 1 1 Dendrocolaptes picumnus 0 0 00 0 0 0 0 1 1 0 0 00 0 0 0 0 0 0 0 0 0 XXI 1X0 0 111 Dendroka graciae 0 0 1 0 10X0X1 XXI 1111X1 X X X X 1X1 110 0 0 0 0 Dendrortyx barbatus Oil 0 0 0 0 0 0 0 0 0 0o 0 0 0 o o 0 0 0 0 0 0 00 0 00 0 0 0 Dendrortyx leucophrys 0 0 0O 0 0 O O 1 X 0 0 0O 0 O O X X 0 0 0 0 1 1 X1 1 O0 110 Dendrortyx macroura 0 0 10 11110 0 O 1 X10 10 0 0 0 10 0 0 0 00 0 00 0 0 0 Diglossa baritula 0X1 0 1 1 1 1 1 X 0 1 X1111X1 1 1 1 X 1 X X 10 0 0 0 0 0 Diglossa plurnbea 0 0 00000000 o o 00 0 0 0 0 0 0 0 0 0 0 0 00 0 1 1111 Doridia eliza 0 0 10 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 00 0 00 0 0 0 Dorkha enicura 0 0 0O O 0 0 O X X 0 0 0O 0 0 O X X 0 0 0 0 1 X X 10 0 0 0 0 0 Doryfera ludovkiae 0 0 00 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 00 0 01111 Dryocopus lineatus 1 1 X1 O 1 O X 1 X XXI X X X 1 X X 0 0 0 0 1 X X 111 1111 Dysithamnus mentalis 0 0 0O O 0 O O 0 X 0 0 00 0 0 0 0 0 0 0 0 0 0 0 010 1 1111

Elaenia frantzii 0 0 0 00 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 1 X X 111 1111 Electron carinatum O 0 O X 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 X X 111 10 0 0 Elvira chionura o o o o 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 00 0 00 0 11 Elvira cupreiceps 0000 000000 0 0 0 000000 0 0 0 0 0 0 00 0 1110 0 Empidonax affinis 0 0 10 0 1 10 0 0 X 1 1 10 10 0 0 X 1 X 1 0 0 00 0 00 0 0 0 Empidonax albigularis 0 111 X X X 1 X X 1 X X X 1 1 o0 10 1 Empidonax atriceps 0 0 0 00 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 00 0 00 111 Empidonax difficilis 1110 X 1 1 1 O 0 1111 X 1 O O O 1 1 1 X 0 0 0000 0000 Empidonax flavescens 0 0 0 10 0 0 0 1 1 o 0 o 00 0 111 0 0 0 0 111 111 1111 Ergaticus ruber 0X10 X 1 1 1 0 O X 1 1 1110 0 0 1 1 1 X 0 0 00 0 00 0 0 0 Ergaticus zrrsicolor 0 0 0 00 0 0 0 1 1 0 o 0 o0 0 0X1 0 0 0 0 111 0 0 0 0 0 0 0 Eubucco bourcierii 0 0 0 00 0 0 0 0 0 0000 0 0 0 0 0 0 0 0 0 0 0 00 0 0 1111 Eugenes fulgens 1110 1 1 1 1 1 X 1111 11111 1 1 1 X 111 1X0 0 111 Eupherusa cyanopJirys 0000 000000 0 0 0 00 10 0 0 0 0 0 0 0 0 00 0 00 0 0 0 Euphentsa exhnia 0 0X0 0 1 0 X X 1 0 0 0 00 O 1 X X 0 0 0 0 XXX 111 1111 Eupherusa nignventris 0000 000000 0 0 0 0o o a o 0 0000 0 0 00 0 00 1 T 1 Eupherusa poliocerca 0 0 0 00 0 0 0 0 0 0 0 0 110 0 0 0 0 0 0 0 0 0 00 0 00 0 0 0 Eupiionia anneae 0 0 0 00 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 00 0 1 1111 Euphoma t'legantissima 1 1 1 X1 1 1 X 1 X X 1 X X 1 1 1 X X X 1 X X 1 1 X1 X X X 1 1 X Euphonia hirundinacea 1111 O 1 O X X X 0 0 0 00 0 0 0 0 0 0 0 0 1X1 111 1111

Formicarius analis 001 101OX1X 000000000 0 0 0 0 0111111111 Formicanus rufipectus 0000000000 000 0 0 0000 0 0 0 0 0000001111 Geothlyph nelsoni 1110X11100 0 0 0 0 1 1 0 0 0 0 10 1 0000000000 Geotn/gon albifacies 011OO1X1XX 000101 1 ix 0 0 0 0 1111100000 Geotrygon chiriquensis 000 0 000000 000000000 0 0 0 0 0000011111 Blanca E. Herndndez-Banos et al. 270

Appendix i (cont.) Mexico HAB GEO East slope West slope TV Belt Central America A B N S

Gcotrygon costarkensis 000 0 0 0 0 0 0 0 0 0 0 0 0 0 000 0 0 0 0 0 0 0 0 0 1 1 1 1 1 Geotrygon carrikeri OOO 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Geotrygon Inwreucii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 Glciucidium gnomci X X X O 0 1 1 O X X X 1 1111 0 X X 1 1 1X X X X 1 0(10 0 0 Glaucidium janiinii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 Glaucidium minutissimum 1 XX 0 0 1 GraHaria guatimnlensis 0 1 1 1 0 0 1 X X X 0 X X 1 11 1 1 X 0 1 0 0 I 1 1 1 X 1 1 1 1 1

X Habia fuscicauda 1 1 1 1 0 1 0 X X X 0 0 0 0 0 0 <> 0 0 (,<> » <> " X 1 1 1 1 1 1 1 Habia rubica Haplospiza rustica 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 1 1 1 1 Harpyhaliaetus solitarius 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 X 1 0 X X 1 1 1 Heliodoxa jacula 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 Heliomaster longirostris 0 0 1 1 0 1 0 X X X 0 0 0 1 0 1 1 X X 0 0 0 0 X 0 0 1 X 1 1 1 1 1

Henicorhina leucophrys 0 1 1 0 0 1 1 1 1 1 0 1 1 1 11 1 1 1 1 0 0 0 I 1 1 1 X 1 1 1 1 1

Henicorhina leitcosticta 0 1 X 1 0 1 0 X X X 0 0 0 0 0 0 I X X 0 0 0 0 I 1 1 1 I ] I 1 1 1

Hylochciris Icucotis 0 1 1 0 1 1 1 1 1 X X 1 1 1 11 1 X 1 1 11 X 1 1 1 1 1 0 0 0 0 0

Hylomanes momotula 0 0 0 1 0 1 0 X X X 0 0 0 0 0 0 X X X 0 0 0 0 1 X X 1 X 1 0 0 0

Icterus graduacauda 1 1X 0 1 1 0 1 0 0 0 1 1 1 11 0 0 0 0 X 0 0 0 0 0 0 0 0 0 0 0 0 Junco phaeonotus X X 1 0 0 1 1 1 1 X 1 X 1 1 1 1 0 X 1 1 10 1 1 1 0 0 0 0 0 0 0 0

Lampornis amethyst inns 1 1 1 1 1 1 1 1 1 X 0 1 1 1 0 1 0 1 1 1 11 X 1 1 1 1 0 0 0 0 0 0 Lampornis calolaemci 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 t) 0 0 0 01 1 1 1 0 Lampornis cinereicauda 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 Lampornis castaneoventns 0 0 0 0 0 0 0 0 0 0 0 0 0 C) 0 00 t) 0 0 0 (} 0 0 0 0 0 0 0 0 0 0 1 Lampornis clemenciae X X 1 0 1 0 1 X 0 0 X X 1 1 1 1 0 0 C) 1 1 1 1 0 (1 00 0 0 0 0 0 0 Lampornis sybdlae 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 Lampornis hemileucus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 X Lampornis viridipallens 0 0 0 0 0 0 0 0 1 X 0 0 0 0 0 0 1 1 1 0 0 0 0 1 T 1 1 0 0 0 0 0 0 Lamprolaima rhami 0 0 X 0 1 1 1 1 1 X 0 0 0 1 0 1 0 1 I 0 X 0 0 1 1 1 1 0 0 0 0 0 0

Lepidocolaptes affin is 1 1 1 1 1 1 1 1 1 1 0 0 0 1 11 1 1 1 0000 1 1 1 1 X 1 1 1 1 1 Lepidocoiaptes leucogaster 0 0 X 0 0 1 1 0 0 0 X 1 1 X 1 1 0 0 0 X 1 1 0 0 C) 0 0 0 0 0 0 0 0

Leptopogon amauroccphalus 0 0 0 1 0 1 0 X X X 0 0 0 0 0 0 0 0 0 0 0 0 0 X X 1 1 X I 1 1 X 1

Leptopogon superciliaris 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 Leptotda z'erreauxi 1 1 1 1 X X 1 X X 1 X X 1 X X X 1 1 X X X 1 X 1 X X 1 1 1 1 1 1 1

Leucopternis albicollis 0 0 X 1 0 1 0 1 X X 0 0 0 0 0 0 X X X 0 0 0 0 X X X 1 1 1 1 1 1 1

Leitcopternis princeps 000 0 0 0 0000 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 Lophostrix cristata

Lophotriccus pileatus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 > 1 1 1 Loxiti curvirostra X X ,\ u 0 1" Lysurus crassirostris 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1

Margaronus rubigmosus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 I 1 1 1 Megarynchus pitangua 1 11 1 X X 1 1 1 X X X X X I X X 1 1 1 1 1 1 Melanerpes formkivorus 1 11 0 1 1 J 1 1 X X I 1 1 1 I 1 1 X 1 1 X 1 1 1 1 1 X I 1 1 1 Melanotis caerulescens 1 11 0 1 1 1 1 0 0 1 1 1 1 11 0 0 0 1 I1 X 0 0 0 0 0 0 0 0 0 0

Melanotis hypoleucus 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 1 X 0 0 0 0 0 0

Micrastur ruficollis 0 1 1 1 0 1 0 X X X 0 0 1 0 0 1 1 X X 0 0 0 (} X X 1 1 X 1 1 1 1 1

Microcerculus marginatus 0 0 0 0 0 0 0 0 X 1 0 0 0 0 C) 0 0 0 0 t) 0 (1 X X X 1 X 1 1 1 X 1

Mionectes oleagineus 0 0 1 ] 0 1 0 X X X 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 X 1 X 1 1 1 1 X

Aiitrephanes phaeocercus 1 11 0 1 1 1 1 1 X "1 11 "1 I1 X I X 1 :1 X 1 1 1 1 1 1 1 ] 1 1

Momotus momota 1 1 1 1 0 1 0 X X X 0 0 0 000 1 X X 0 0 0 0 I 1 I 1 1 1 1 ] 1 I

Myadestes melanops 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1

Myadestes occidental is 1 11 0 1 1 1 1 1 X 1 11 1 11 1 1 1 1 11 X 1 1 1 I 0 0 0 0 0 0

Myadestes unicolor 0 I 1 ^ 0 1 0 1 1 1 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 1 1 1 0 0 0 0 0

Myiarchus tuber cidifer 1 11 1 0 1 1 X 1 X X X 1 1 1 1 1 X X 1 I 1 X ] 1 1 1 1 1 1 1 1 X

Mywbius su!phurep\/gius 0 0 1 1 0 1 0 X X X 0 a 0 0 0 0 0 0 0 C) 0 X X 1 X 1 1 1 1 1 1 Myioborus miniatus 0 1 1 1 1 1 1 1 1 1 X 1 1 1 11 1 1 1 1 1 1 X 1 1 1 1 X 1 1 1 1 1

Myioborus pktus 1 11 X X X X 1 1 X 1 1 1 XX X 1 1 1 1 1 X 1 1 1 1 1 0 0 0 0

Myioborus torquatus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (1 00 0 1 1 1 1

Myicdynnstes hemkhryseus 0 0 0 0 0 0 0 Q 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 Bird faunas of Mesoamerican humid montane forests 271

Appendix 1 (cont.) Mexico HAB GEO East slope West slope TV Belt Central America A B N S

Myiodynastes luteiventris 111X0X0XXX XXXXXX1XX 0 0 0 0 1XX1X111 0 0 Myiopagis viridicata XX11X1XXXX XI 1 1 X X X XX X 1 1 X 1 X X 1 1 1 1 11 1 Myrmeciza immaculata 00000 0 0000 0 0 0 0 0 0 0 00 0 0 0 0 000001 1 1 1 1 Myrmotherula schisticolor 00000000X1 0000000X1 0 0 0 0 0XX1X1 1 11 1 Nothocercus bonapartei ooooocoooo 000000000 0 0 0 0 00000101 1 1 Nyctidromus albkoilis 1111XXXXXX X 1 X X X X 1 IX X 0 0 0 XX111111 1 1 Nyctiphrynus mcieodii 0000000000 010001000 0 0 0 0 0000000000 Odontophorus guttatus OOX 1 OXOXXX 0000001 XX 0 0 0 0 1XX11XX1 1 1 Odontophorus leucolaemus 0 0 0 0 0 0 0 0 0 0 000000000 0 0 0 0 000001 1 11 1 Oreophasis derbianus 00 0 0000000 000000011 0 0 0 0 1000000000 Otus barbarus ooooooooxx 000000000 0 0 0 0 01 00000000 Otus choliba 0000000000 000000000 0 0 0 0 0001XX1 1 1 1 Otus darkii 0000000000 000000000 0 0 0 0 000001 1 11 1 Otus flammeolus 0010 0 00000 01 1 100000 X 1 1 X xooooooooo Otus guatemalae 1X1XXXXX1X xxooooooo 0 0 0 0 0 X X 1 X 1 1 11 1 Otus trichopsis XX10001000 0X1X110 00 1 1X10000 0 0 Pachyramphus aglaiae 1 1 X X Pachyramphus albogriseus 1111XX1XXX X1XXXX11X X 1 1 X 1 XXXX1 0000 Pachyramphus major 0000000000 000000000 0 0 0 0 0000001 1 1 1 Pachyramphus verskolor 1111010XXX 11110 1111 0 0 0 0 1111100000 Panterpe insignis 0000000000 000000000 0 0 0 0 0 0 0 0 0 0 1 11 1 Panyptila sanctihieronymi 0000000000 000000000 0 0 0 0 00000111 1 1 Parula gutturalis 0000000000 001X11110 X 0 0 0 1XX1000000 Parula pitiayumi 0000000000 000000000 0 0 0 0 00000001 1 1 Parula superciliosa 1 100000000 XX01 00010 0 0 0 0 1X11X111 1 1 Parus sclateri 111011111X 111111X11 1 1 1 X 111110000 0 Penelope purpurascens 0010011000 XXI 101 000 11X1 000000000 0 Penelopina nigra 11X101011X 11110111X 0 0 0 0 XXX1X1 1 1 1 1 Peucedramus taeniatus 000000001X 0000001 11 0 0 0 0 11X1100000 Pezopetes capitalis 1X101110XX 1X1 1010XX 1 X X 1 1111100000 Phaethornis guy 0000000000 000000000 0 0 0 0 00000001 1 1 Phaethornis longuemareus 0000000000 000000000 0 0 0 0 000001 1 1 1 1 Phaethornis superciliosus 001 1010XXX 000000000 0 0 0 0 XXX111I1 1 1 Phainoptila melanoxantha 00X1010XXX 01X1 1 1XXX 0 0 0 0 XXXX1111 1 1 Pharomachrus mocinno 0000000000 000000000 0 0 0 0 00000111 1 1 Pheucticus chrysopeplus 000000001 1 0000001 IX 0 0 0 0 11111011 1 1 Pheucticus melanocephalus 1XX00000 Pheucticus tibialis 11X0011X00 XX11 11000 1 1 1 X 00000000 0 0 Philydor rufus 0000000000 000000000 0 0 0 0 000001 1 1 1 1 Phyllomyias burmeisteri 0000000000 000000000 0 0 0 0 0000001 1 1 1 Phylloscartes superciliaris 0000000000 000000000 0 0 0 0 0 0 0 0 0 0 1 11 1 Piaya cayana 0000000000 000000000 0 0 0 0 00000111 X 1 Picoides villosus 1111XXXXXX XXI 1XX1XX 1 1 1 X Piculus auricularis 1 1 Piculus rubiginosus 0000 0 0 0000 11X111000 0 0 0 0 00000000 0 0 Pionopsitta haematotis 11X1X1XX1X ooooooixx 0 0 0 0 11111111 1 1 Pionus senilis 0001010100 000000000 0 0 0 0 0XX11111 I 1 Pipilo erythrophthalmus 1 1XX0X0XXX OOOOOOIXX 0 0 0 0 0X111111 1 1 Pipilo ocai 1110111111 1110110X1 0 111 11000000 0 0 Pipra pipra 0010111100 01X111000 10 0 0 0000000000 Piranga bidentata 0000000000 000000000 0 0 0 0 00000001 1 1 Piranga erythrocephala 1X1001001X 011111111 1 1 1 X 1111XXX1 1 1 Piranga flava 00000X1 1 0 0 111111000 X 1 X X 0000000000 Piranga leucoptera XX1X1 1 1 1XX 1X111111X 1 1 1 X XI111111 1 1 Platyrinchus cancrominus 1I11010XXX 0000001 IX 0 0 0 0 1XX11111 1 1 Platyrinchus mystaceus 0OX1O10XXX 000000000 0 0 0 0 1XX1 1 10000 Premnoplex brunnescens 0000000000 000000000 0 0 0 0 000001 1 11 1 Procnias tricarunculata 0000000000 000000000 0 0 0 0 000001 1 I1 1 Pselliophorus tibialis 0000000000 000000000 0 0 0 0 00011111 1 1 0000000000 000000000 0 0 0 0 00000011 1 1 Blanca E. Herndndez-Banos et al.

Appendix 1 (cont.) Mexico HAB GEO East slope West slope TV Belt Central America A B NS

PselHophorus luteoviridis o0 ooo 0000 0 o 0 o a a o o o o 0 0 0 0 0000000001 Pseudocolaptes lawrencii 0000 000000 0 0 0 0 0 0 0 0 0 0 0 0 0 0000001111 Ptilogonys caudatus 0000000000 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0000001 1 1 Ptilogonys cinereus 111O11111X 1X111 1 0 1 X 1 1 1 X 1 1X0000000 Pyrrhura hofpnanni 0000000000 0 0 0 0 0 0 0 0 0 0 0 0 0 0000000 0 1 1 Ramphocelus saiiguinoleiitis 0011010XXX 0 0 0 0 00 0 0 0 0 a o o XXXX111111 Regulus satrapa OOOOOO1OXX 0 0X10 0 0X1 1 X X X X1X0000000 Rhynchocyclus brevirostris 0 0 1 1 0 1 O O X X 0 0 0 1 11 1 X X 0 0 0 0 1111111111 Rhynchopsitta pachyrhyncha 0000000000 X X 0 0 O 0 0 0 0 1000 0000000000 Rhynchopsitta terrisi 1 000000000 0 0 0 0 00 0 0 0 o o o o 0000000000 Ridgzvayia pinicola 0X10XX1X00 X 1 1 1 X X X X X 00 00000000 Sayornis nigricans 1 1 1XXXXXX1 1 Sclerurus albigularis 1111111111 0000000000 0 0 0 0 00 0 0 0 0 0 0 0 Sderurus mexicanus 0000001 1 1 1 001X0100XX 0 0 0 0 00 110 0 0 0 0 Scytalopus argentifrons 1111X11111 0000000000 0 0 0 0 00 0 0 0 0 0 0 0 Selasphorus ardens 000001 1 1 1 1 0000 0 00000 0 0 0 0 00 0 0 0 0 0 0 0 Selasphorus flammula 0000000001 0000000000 0 0 0 0 00000 0 0 0 0 Selasphorus platycercus 0O0O0O01 1 1 Selasphorus scintilla 1 XXOOOOOOO 000000000 0 0 0 0 0 000001111 Semnornis frantzii 0 0 0 0 0 0 0 0 0 0 0000000000 Serpophaga cinerea 0 0 0 0 0 0 0 0 0 0 0 0 0 0000001 1 1 1 0000000000 0 0 0 0 0 0 o o o o 0000001111 Sitta carolinensis 0 0 0 XXX010100 0 1110 0 0 X X X X 0 0 0 0 0 0 0 0 0 0 Sitta pygmaea XXI X X 1 O 0 0 O 0 0 O 0 0 0 0 0 0 X X X X Sittasotnus griseicapillus XXI 0 0 0 0 0 0 0 0 0 0 Spizaetus oruatus 1 1X101001X Oil 10 1 1 X X 0 0 0 0 1X11X11111 Streptoprocne semicollaris 1XX1X1XXXX 0 0 0 10 0 0 0 0 0 0 0 0 XXX1X11111 Streptoprocne zonaris 0000000000 XXI 100000 X X O 0 0000000000 Strix fulvescens 111101111X 0 0 0 X 1 1 1 X 1 X 0 1 0 111111X111 Strix varia 000000001 X 0 0 0 0 0 0 0 1 1 0 0 0 0 1111000000 Syndactyla subalaris OXX0001000 O X X 1 0 0 0 0 0 0 0 0 0 0000000000 Tangara cabanisi 0000000000 0 0 0 0 0 0 0 0 0 0 0 0 0 000001 1 1 1 1 Tangara down 0000000000 0 0 00 0 0 0 1 1 0 0 0 0 1000000000 Tangara gyrola 0000000000 0 0 00 0 0 0 0 0 0 0 0 0 0000001111 Tangara icterocephala 0000000000 0 o 0000000 0 0 0 0 0000001111 Terenura callinota 0000000000 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 Thalurania ridgwayi 0000000000 0 0 0 0 0 0 0 0 0 0 0 0 0 0000001 1 1 1 Thamnophihts doliatus 0000000000 010 000000 0 0 0 0 0000000000 Thraupis abbas 1111010XX1 0 0 0 0 0 0 1 IX 0 0 0 0 1XX1X11111 Thripadectes rufobrunneus 1111010X1 X 0 0 0 0 0 0 1X1 o 0 o o 1X11100000 Thryothorus maculipcctus 0000000000 0 0 0 0 0 0 0 0 0 0 0 0 0 0000011111 Thryothorus modestus 111101011X 000 0 O O O X X 0 0 0 0 1X11100000 Thryothorus rufalbus 0000000000 0 0 00 O O X X X 0 0 0 0 1XX1X11111 Tityra semifasciata 0000000000 0 0 00 0 0 0 1 1 0 0 0 0 xxxxxooooo Tohnomyias sulphurescens 1 1 1 1 O 1 0 X X X XXX 1X1 1 X X 0 0 0 0 1111111111 Touit costaricensis 0011010XXX 0 0 0 O O 0 1 X X 0000 1X11X11111 Toxostoma ocellatum 0000000000 0 0 0 0 0 0 0 0 0 0 0 0 0 00000001 1 1 Troglodytes brunnekollis 0010001000 0 0 0 0 1 0 0 0 0 0 X 1 X 0 000000000 Troglodytes ochraceus X111111100 XXI 1110 0 0 1 1 1 X 0000000000 Troglodytes rufociliatus 0000000000 0 0 0 0 0 0 0 0 0 0 0 0 0 0000001111 Trogon aurantiiventris 000000001X 0 0 0 O 0 O O I 1 0 0 0 0 1 1 1 1 100000 Trogon collaris 0000000000 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 Trogon elegans 01110100XX 0 0 0 1 1 1 1 1 X 0 0 0 0 1X11100111 Trogon mexicanus 11X00001XX 111 X O 1 0 1 X X X 0 0 1X1 1X10000 Turdus assimilis 111011111X 111 1 1 1 0 1 X 1110 1111000O0O Turdus grayi 11110111X1 1 1 1110 1XX1111111 Turdus infuscatus 111101001X o 0 o O O O 1 1 X 0X00 1X1111111X Turdus migratorius 1 1 1 1 0 1 1 1 1 X 0 0 0 111111 0 0 0 0 1111000000 Turdus nigrescens X110111100 XXI 1110 0 0 X 1 X 1 0000000000 Turdus pkbejus 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0000000111 0000000000 0 0 00 0 0 11 1 o o o o 1 1 1 1 1 1 I I 1 1 Bird faunas of Mesoamerican humid montane forests 273

Appendix 1 (cont.) Mexico HAB GEO East slope West slope TV Belt Central America A B N s

Tunius nifitoraues 0 0 0 0 OOOOOOOOI 1 O00OO0 0X1 0000 1111 0 0 0 0 0 0 Tyto alba 1 1 1 1 XXllXXXXXX 1 X 1 XX 1 1 1 1 I

Ve liliot'ms futnigatus 0 1 0 1 1 1X1X1XXXX (XXIX 0X00 XXXlllllI 1 Vir eo breuipewiis 0 1 0 0 OOIOOOOXOO 0100 10 0 0 0 0 10 000000000 0 Vir eo carmioti 0 0 0 0 oooooooooo 000000000 0 0 0 0 000000 0 1 1 1 Vir eo huttoni 1 0 1 0 XX1O111111 1111 1X11 1 X 1 X 1110 0 0 0 0 0 0 Vir eo leucophrys 0 0 0 1 1 1 1OOXOX1X OOOOI ) 0 1 X X 0 0 0 0 XXXI X 0 1 1 1 1 Vi eo solitarius 1 0 1 0 XX1O1O1XXX 0X11 1 1 X X 1 1 1 X 1111 0000 0 0 Vi eolntiius tnelitophrys 0 1 0 0 01 10001XXX 0 111 1 0 x x 1110 10 0000000 0

Xii hocolaptes f romeropirhynchus 0 1 0 1 0 1XX011111 000101000 0 0 0 0 0 X X 1 1 X X 1 1 1 horhynchus erythropygius 0 0 0 1 OIXOOIOXIX OOOIOOIIX 0 0 0 0 1X11 11111 1

Zeledonin coronata 0 0 0 0 oooooooooo 000000000 0000 0 0 0 0 01111 1 Zimmerius vilissimus 0 1 0 1 0000000 0 XX 00000001 X 0 0 0 0 1111 X 1 1 1 1 X

Appendix 2. Descriptions of the 33 humid montane forest patches included in this study. HPO, humid pine-oak forest; CF, cloudforest. General references are listed in the Methods section; here listed as references are major sources that provided numerous records for particular islands. Abbreviation Name Habitat References Eastern Mexico 1. TAMPS Tamaulipas and Nuevo Leon HPO, CF Harrell (1951) 2. HIDAL N Veracruz, Hidalgo, SLP HPO, CF Sutton and Burleigh (1940), Lowery and Newman (1949) 3. CVERA Central Veracruz HPO, CF Chapman (1898), Navarro-Sigiienza et al. (1991) 4. TUXTL Los Tuxtlas, Veracruz CF Wetmore (1943), Andrle (1967) 5. CUATR Sierra de Cuatro Venados, Oaxaca HPO Binford (1989), Rowley (1984) 6. JUARE Sierra de Juarez, Oaxaca HPO, CF Binford (1989), Torres-Chavez (1992) 7. FELIP Cerro San Felipe, Oaxaca HPO Binford (1989) 8. ZEMPO Cerro de Zempoaltepetl, Oaxaca HPO, CF Binford (1989) 9. SCRIS San Cristobal de las Casas, Chiapas HPO, CF Alvarez del Toro (1980) 10. MONTE Lagunas de Montebello, Chiapas HPO, CF Alvarez del Toro (1980) Western Mexico 11. SINDU Sinaloa and Durango HPO 12. NAYJA Nayarit and Jalisco HPO Escalante-Pliego (1988), Schaldach (1963) 13. MICHO Coastal Michoacan HPO — 14. GUERR Sierra de Atoyac, Guerrero HPO, CF Navarro-Sigiienza (1986, 1992) 15. YUCUN Sierra de Yucunacua, Oaxaca HPO Binford (1989) Blanca E. Herndndez-Banos et al. 274

Appendix 2 - (cont.) Abbreviation Name Habitat References 16. MIAHU Sierra de Miahuatlan, Oaxaca HPO Binford (1989), Rowley (1966) 17. CHIMA Los Chimalapas, Oaxaca HPO, CF Binford (1989) 18. SOCON Sierra de Soconusco, Chiapas HPO, CF Parker et al. (1976), Alvarez del Toro (1980) 19. TACAN Volcan de Tacan HPO, CF Alvarez del Toro (1980) Transvolcanic Belt, Mexico 20. TANCI Cerro de Tancitaro, Michoacan HPO, CF Blake and Hanson (1942) 21. TEMAS Temascaltepec, Mexico HPO Ornelas et al. (1988) 22. TAxCO Sierra de Taxco, Guerrero HPO Morales-Perez and Navarro-Sigiienza (1992) 23. MALIN Volcan La Malinche HPO Central America 24. GUAPA Pacific slope, Guatemala HPO, CF Griscom (1932), Land and Wolf (1961) 25. GUACU Sierra de los Cuchumatanes, HPO, CF Griscom (1932), Land and Guatemala Wolf (1961) 26. GUAMI Sierra de las Minas, Guatemala HPO, CF Griscom (1932), Land and Wolf (1961) 27. HONDU Honduras and El Salvador HPO, CF Monroe (1968), Thurber et al. (1987) 28. NICAR Nicaragua HPO, CF Slud (1964), Stiles and 29. CRGUA Cordillera de Guanacaste, Costa Rica HPO, CF Skutch (1989) Slud (1964), Stiles and 30. CRTIL Cordillera de Tilaran, Costa Rica HPO, CF Skutch (1989) Slud (1964), Stiles and 31. CRCEN Cordillera Central, Costa Rica HPO, CF Skutch (1989) Slud (1964), Stiles and 32. CRTAL Cordillera de Talamanca HPO, CF Skutch (1989) Wetmore (1965-1984) 33. WPANA Western Panama HPO, CF

References Alvarez del Toro, M. (1980) Las aves de Chiapas. Tuxtla Gutierrez, Chiapas: Gobierno del Estado de Chiapas. Andrle, R. F. (1967) Birds of the Sierra de Tuxtla in Veracruz, Mexico. Wilson Bull. 79: 163-187. AOU (1983) Check-list of North American birds. Washington, D.C.: American Ornitholo- gists' Union. Banks, R. C. (1990) Taxonomic status of the Coquette hummingbird of Guerrero, Mexico. Auk 107: 191-192. Baptista, L. F. (1978) A revision of the Mexican Piculus (Picidae) complex. Wilson Bull. 90: 159-181. Binford, L. C. (1989) A distributional survey of the birds of the Mexican state of Oaxaca. Orn. Monogr. 43: 1-405. Blake, E. R. and Hanson, H. C. (1942) Notes on a collection of birds from Michoacan, Mexico. Field Mus. Nat. Hist., Zool. Ser. 22: 513-551. Chapman, F. M. (1898) Notes on birds observed at Jalapa and Las Vigas, Vera Cruz, Mexico. Bull. Amer. Mus. Nat. Hist. 10: 15-43. Bird faunas of Mesoamerican humid montane forests 275

Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Naranjo, L. G., Parker, T. A. and Wege, D. C. (1992) Threatened birds of the : the ICBP/IUCN Red Data Book. Cambridge, U.K.: International Council for Bird Preservation. Cracraft, J. (1983) Species concepts and speciation analysis. Curr. Orn. 1: 159-187. Escalante-Pliego, P. (1988) Las aves de Nayarit. Mexico: Universidad Autonoma de Nayarit. Escalante-Pliego, P. and Peterson, A. T. (1992) Geographic variation and species limits in Middle American woodnymphs (Thalurania). Wilson Bull. 104: 205-219. Escalante-Pliego, P., Navarro-Sigiienza, A. G. and Peterson, A. T. (1993) A geographic, ecological, and historical analysis of land bird diversity in Mexico. Pp.281-307 in T. P. Ramamoorthy, R. Bye, A. Lot and J. Fa, eds. Biological diversity of Mexico: origins and distribution. New York: Oxford University Press. Friedmann, H., Griscom, L. and Moore, R. T. (1950) Distributional check-list of the birds of Mexico, Part 1. Pacific Coast Avifauna 29: 1-202. Graham, A. (1993) Historical factors and biological diversity in Mexico. Pp. 109-127 in T. P. Ramamoorthy, R. Bye, A. Lot and J. Fa, eds. Biological diversity of Mexico: origins and distribution. New York: Oxford University Press. Griscom, L. (1932) The distribution of bird-life in Guatemala. Bull. Amer. Mus. Nat. Hist. 9: 1-439. Harrell, B. E. (1951) The birds of Rancho del Cielo, an ecological investigation in the oak- sweet gum forest of Tamaulipas, Mexico. Thesis, University of Minnesota. ICBP (1992) Putting biodiversity on the map: priority areas for global conservation. Cambridge, U.K.: International Council for Bird Preservation. Land, H. C. and Wolf, L. L. (1961) Additions to the Guatemalan bird list. Auk 78: 94-95. Long, A. J. (1995) The importance of tropical montane cloud forests for endemic and threatened birds. Pp.79-105 in J. O. Juvik and F. N. Scatena, eds. Tropical montane cloud forests. New York: Springer-Verlag. Lowery, G. H., Jr. and Newman, R. J. (1949) New birds from the state of San Luis Potosi and the Tuxtla Mountains of Veracruz, Mexico. Louisiana State Univ. Mus., Occas. Pap. 22: 1-10. Martin, P. S. and Harrell, B. E. (1959) The Pleistocene history of temperate biotas in Mexico and eastern United States. Ecology 38: 468-480. McKitrick, M. C. and Zink, R. M. (1988) Species concepts in ornithology. Condor 90: 1-14. Miller, A. H., Friedmann, H., Griscom, L. and Moore, R. T. (1957) Distributional check- list of the birds of Mexico, Part 2. Pacific Coast Avifauna 33: 1-436. Monroe, B. L., Jr. (1968) A distributional survey of the birds of Honduras. Orn. Monogr. 7: 1-458. Morales-Perez, J. E. and Navarro-Sigiienza, A. G. (1992) Analisis de la distribucion de las aves en la Sierra Norte del Estado de Guerrero, Mexico. An. hist. Biologia, U.N.A.M. 62: 497~51°. Navarro-Siguenza, A. G. (1986) Distribucion altitudinal de las aves en la Sierra de Atoyac, Guerrero. Tesis profesional, Facultad de Ciencias, U.N.A.M. Navarro-Sigiienza, A. G. (1992) Altitudinal distribution of birds in the Sierra Madre del Sur, Guerrero, Mexico. Condor 94: 29-39. Navarro-Sigtienza, A. G., Morales-Perez, J. E. and Hernandez-Banos, B. E. (1991) Aves de Teocelo. Teocelo 7 pp. Navarro-Sigiienza, A. G., Peterson, A. T., Escalante-Pliego, P. and Benitez-Diaz, H. (1992) Cypseloides storcri, a new species of swift from Mexico. Wilson Bull. 104: 55-64. Ornelas, J. F., Navarijo, L. and Chavez, N. (1988) Analisis avifaunistico de la localidad de Temascaltepec, Estado de Mexico, Mexico. An. hist. Biologia, U.N.A.M. 58: 373- 388. Parker, T. A., Ill, Hilty, S. and Robbins, M. B. (1976) Birds of El Triunfo cloud forest, Mexico, with notes on the Horned Guan and other species. Amer. Birds 30: 779-782. Blanca E. Herndndez-Banos et al. 276

Peterson, A. T. (1992) Phylogeny and rates of molecular evolution in the Aphelocoma jays (). Auk 109: 133-147. Peterson, A. T. (1993) Species status of Geotrygon carrikeri. Bull. Brit. Orn. Club 113: 166- 168. Peterson, A. T., Escalante-Pliego, P. and Navarro-Siglienza, A. G. (1992) Genetic vari- ation and differentiation in Mexican populations of Common Bush-tanagers and Chest- nut-capped Brush-finches. Condor 94: 244-253. Peterson, A. T., Flores-Villela, O. A., Leon-Paniagua, L. S., Llorente-Bousquets, J. E., Luis-Martmez, M. A., Navarro-Sigiienza, A. G., Torres-Chavez, M. G. and Vargas- Fernandez, I. (1993) Conservation priorities in northern Middle America: moving up in the world. Biodiversity Letters 1: 33-38. Ramamoorthy, T. P., Bye, R., Lot, A. and Fa, J. (1993) Biological diversity of Mexico: origins and distribution. New York: Oxford University Press. Remsen, J. V. (1994) Use and misuse of bird lists in community and conservation. Auk 111: 225-227. Rowley, J. S. (1966) Breeding records of birds in the Sierra Madre del Sur, Oaxaca, Mexico. Proc. Western Found. Vert. Zool. 1: 107-204. Rowley, J. S. (1984) Breeding records of land birds in Oaxaca, Mexico. Proc. Western Found. Vert. Zool. 2: 73-224. Rzedowski, J. (1986) Vegetacion de Mexico. Mexico City: Editorial Limusa. Sarukhan, J. and Dirzo, R. (1992) Mexico confronts the challenges of biodiversity. Mexico City: Comision Nacional para el Conocimiento y Uso de la Biodiversidad. Schaldach, W. J., Jr. (1963) The avifauna of Colima and adjacent Jalisco, Mexico. Proc. Western Found. Vert. Zool. 1: 1-100. Slud, P. (1964) The birds of Costa Rica: distribution and ecology. New York: American Museum of Natural History. Soule, M. E. (1990) The real work of systematics. Ann. Missouri Bot. Garden yy: 4-12. Stiles, F. G. and Skutch, A. F. (1989) A guide to the birds of Costa Rica. Ithaca, New York: Cornell University Press. Sutton, G. M. and Burleigh, T. D. (1940) Birds of Valles, San Luis Potosi, Mexico. Condor 42: 259-262. Torres-Chavez, M. G. (1992) Distribution altitudinal de las aves en la Sierra de Juarez, Oaxaca. Tesis profesional, Facultad de Ciencias, U.N.A.M. Thurber, W. A., Serrano, J. F., Sermeno, A., Benitez, M. (1987) Status of uncommon and previously unreported birds of El Salvador. Proc. Western Found. Vert. Zool. 3: 109- 293. Vazquez-Garcia, J. A. (1995) Cloud forest archipelagoes: preservation of fragmented montane ecosystems in tropical America. Pp.315-332 in J. O. Juvik and F. N. Scatena, eds. Tropical montane cloud forests. New York: Springer-Verlag. Wetmore, A. (1943) The birds of southern Veracruz, Mexico. Proc. U.S. Natn. Mus. 93: 215-340. Wetmore, A. (1965-1984) The birds of the Republic of Panama, Parts 1-4. Washington, D.C.: Smithsonian Institution.

BLANCA E. HERNANDEZ-BANOS Museo de Zoologia, Facultad de Ciencias, Universidad Nacional Autonoma de Mexico, Apartado Postal 70-399, Mexico, D.F. 04510, Mexico

A. TOWNSEND PETERSON Natural History Museum, The University of Kansas, Lawrence, Kansas 66045, U.S.A. Bird faunas of Mesoamerican humid montane forests 277

ADOLFO G. NAVARRO-SIGtJENZA Museo de Zoologia, Facultad de Ciencias, Universidad National Autdnoma de Mexico, Apartado Postal yo-399, Mexico, D.F. 04510, Mexico

PATRICIA ESCALANTE-PLIEGO Departametito de Zoologia, Instituto de Biologia, Universidad National Autdnoma de Mexico, Apartado Postal 70-153, Mexico, D.F. 04510, Mexico